Sexual conflict between two conspecific sexes arises when divergent interests about the timing and duration of copulation, number of mating partners, or parental investment occur, provided that a trait enhancing the reproductive success (~fitness) of one sex may reduce that of the other (Arnqvist and Rowe, 2005). Currently more broadly defined as the sexually antagonistic selection of shared traits, the sexual conflict is now recognised as a “pervasive evolutionary selective force“, leading in many known instances to a male-female coevolution (Parker, 1979; Rostant et al., 2020). At the very core of this conflict lies the struggle for dominance over genetic control in offspring by each sex (Thornhill, 1976; Thornhill, 1979). The study of mating practices offers one of the most direct ways to address sexual conflict in nature. Two seemingly opposite (yet not necessarily mutually exclusive) mating-related strategies are observed in males: aggressive behaviour toward a female, which allows exerting domination while mating, and the nuptial gifting of food to allure and/or pacify a female during the time of copulation and sperm transfer (Lehmann, 2012; McCartney et al., 2013). Both coercive and nuptial gifting strategies appear costly to males because they are based on the differential allocation of limited resources (Liu et al., 2015); therefore, an exaggerated armature may hinder the overall fitness in a male (e.g. in rhinoceros and flour beetles a larger ‘weapon’ is negatively correlated with wing size; Yamane et al., 2010). In order to incline the female to mate, males show aggression (e.g. calopterygid damselflies; Rivera and Andrés, 2002), intimidating behaviour (e.g. water striders; Han and Jablonski, 2010), and may use special structures, such as modified front legs to clasp the female wings (e.g. sepsid flies; Eberhard, 2002) or adhesive discs to secure grip on the female (e.g. diving beetles; Bergsten et al., 2001). Within coercive mating, extreme cases of aggressive behavior by males, which may cause damage or even death to the female, are known among insects and other invertebrate groups, such as true bugs (Siva-Jothy, 2006), water bugs, and camel-spiders (Arnqvist and Rowe, 2002; Hrušková-Martišová et al., 2010) or snails (Kimura and Chiba, 2015), but also among vertebrates (e.g. ducks; Brennan and Prum, 2012). Nuptial gifting is also widespread. Crickets and bush crickets (Orthoptera: Tettigonidea, Grylloidea) provide an edible gift attached to a structure containing the sperms. Female consume the nutritious part during fertilisation by the spermatozoans, the time for consumption correlating with that of fertilisation (Lehmann, 2012; McCartney et al., 2013). Males of other orthopterans offer thoracic secretions (Gryllidae; Bussière et al., 2005) or even their own hind wings as nuptial gifts (Haglidae; Eggert and Sakaluk, 1994; Sakaluk et al., 2004). Silk-wrapped nuptial gifts of prey are offered by spiders (Stålhandske, 2001) and dance flies (Diptera: Empididae; Preston-Mafham, 1999; Sadowski et al., 1999; Lebas and Hockham, 2005); the shining silk cover makes a gift more enticing and may deceive the female if it only contains a scarce – or even fully absent – edible portion (Ghislandi et al., 2017). In dobson flies (Megaloptera: Corydalidae), the species offering nuptial gifts have a ‘weapon’ armature used for fighting off the rivals smaller than in species whose males rely only (or mostly) on such armature (Liu et al., 2015). Among mecopterans, a gift of prey is offered by hangingflies (Bittacidae; Thornhill, 1976) and scorpionflies (Panorpa spp.; Gwynne, 1984); the latter may offer hardened salivary masses to the female instead (Thornhill, 1979; Bockwinkel and Sauer, 1994). Coercive and nuptial-gifting mating approaches can co-occur in same species, which then exhibits a mixed strategy that can serve as a model to study the trade-off between these two tactics (e.g. Tong and Hua, 2019).

Scorpionflies (Mecoptera: Panorpidae) are models in sexual conflict research. Their mating practices range from coercive to provisioning nuptial gifts, in extreme cases including mouth-to-mouth salivary transfer until the end of copulation (Zhong et al., 2015). Mecoptera are among the oldest holometabolous insects. They are known since the Permian, when they were a significant component of the flying insect fauna. Whereas scorpionflies flourished in the Mesozoic, most of their lineages rapidly declined during the Cenozoic. In fact, mecopterans are considered to have gone through a degree of extinction unparalleled among holometabolans (Grimaldi and Engel, 2005). Extant Mecoptera are classified in nine families, most of which are relict; only two lineages, Panorpidae and Bittacidae, are relatively species-rich. The Panorpidae appeared in the Late Jurassic (Ding et al., 2014) and currently represent the most diverse mecopteran lineage, with about 480 described species classified in seven genera (Bicha, 2018; Miao et al., 2019), that is, c. 67% of extant mecopteran species.

In scorpionfies, the male behaviour towards the female may be coercive in such a degree that the expression ‘rape’ has been used in the past for describing their mating rituals (Thornhill, 1980). In any case, the role of a female in controlling mating and the eventual reproductive outcome appears more active and decisive than initially thought in the group, namely through precopulatory male choice, struggling behaviour to interrupt genital connection, and the selective reception of sperm from gift-providing males (likely through spermathecal muscular action; Kock et al., 2009). The modifications of panorpid males enabling coercive mating behaviour include a diverse array of structures on different abdominal segments as well as thinned and elongate last segments of the abdomen. Some of these structures, the so-called notal and postnotal organs, effectively act as a ‘clamp’, grasping the female’s wing prior to and during the course of copulation (Figure 1A, B, F). Depending on the size and shape of these organs in a species, the grasp is more or less coercive. Elongate abdominal segments enable the males to bend the abdomen over that of the female, while additional, usually paired abdominal structures referred to as anal horns are used to achieve and maintain the strained, twisted copulatory position of both abdomens for the long time required for sperm transmission, which could be more than 3 hr (Tong et al., 2018; Tong and Hua, 2019). Panorpid species with very long notal organs, capable of clamping the entire width of the female wing, only use coercion, so no food gifts are offered (Tong and Hua, 2019). In species with mixed strategies, it is the physical condition of the male which determines the followed strategy (Engqvist and Sauer, 2001): older and weaker males, as well as smaller ones, primarily adopt an aggressive strategy toward a female, as they are either not able to obtain a prey as a gift or to fabricate it through salivary production, or their gift is often been stolen by a stronger male. Larger males also have an advantage over smaller ones when competing for dead arthropods that can be offered to a female or, if eaten by the male, allow to produce salivary masses. On the other hand, conspecific smaller males bear proportionally larger notal organs (Thornhill, 1990), which shows the genetic background of this negative allometry.

Figure 1

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Here, we describe three exceptional scorpionflies from three different Cretaceous and Eocene ambers deposits and each belonging to a distinct panorpoid lineage. The disparate abdominal configurations and specialisations in the new taxa allow us to extrapolate trends observed among extant relatives, discussing possible mating habits for the new fossils and previously described ones with preserved non-genitalic abdominal modifications (Willmann and Novokschonov, 1998; Krzemiński and Soszyńska-Maj, 2012; Archibald, 2013; Wang and Hua, 2020; Zhang et al., 2021). Additionally, the new taxa are of general evolutionary significance, including the description of a new extinct panorpoid family and the first orthophlebiid species described in amber.

A series of clamping devices used to seize and hold different body parts of the female during courtship and copulation exist in panorpid males. These structures ensure the eventual genital connection and sperm transfer and extend copulation time (Zhong and Hua, 2013a; Zhong et al., 2014). The latter is critical for males, as the more prolonged the copulation, the more sperm can be transferred and, thus, the greater the number of potentially fertilised eggs (Engqvist and Sauer, 2002; Kullmann and Sauer, 2005). Aside from genital clamping structures (among which the most conspicuous is a pair of pincer-like gonostyli), panorpids present non-genital clamping systems on their abdomens. These are notal and postnotal organs, as well as anal horns which are used to catch and immobilize female during mating. Based on the size, shape and interlocking mechanism of these clamping structures, the grasp on the body of female becomes more or less secure, therefore facilitating a greater degree of coercion during mating, which has been associated with less investment in costly nuptial gifts (Tong and Hua, 2019).

Notal and postnotal organs

The notal organ (NO) is a process on abdominal tergite III. The postnotal organ (PO) typically consists of a small process on abdominal tergite IV (Crampton, 1931; Mickoleit, 1971). In extant scorpionflies, both structures are present in most Panorpidae, the only recent species from the Eomeropidae (the ‘living fossil’ Notiothauma reedi MacLachlan, 1877 [MacLachlan, 1877; Crampton, 1931]), and can be vestigial, if present, in the Panorpodidae (Tong et al., 2017; Wang and Hua, 2018a). Both NOs and POs act jointly by typically creating a clamping structure that grasps the female forewing before and during copulation; in panorpids, these organs allow to achieve and maintain the typical V-shaped mating position (Figure 6A and B; Byers and Thornhill, 1983). Moreover, the NO and the PO bear specialised setae, cephalically directed and thinned terminally, which interlace and/or electrostatically interact with both venational setae and membrane microtrichia from the female’s wing (Zhong and Hua, 2013a). Observations on three species of Neopanorpa van der Weele, 1909 (van der Weele, 1909) bearing NOs of different lengths have shown different mating strategies in these species (Zhong and Hua, 2013a; Tong and Hua, 2019). Males of two species with short and medium-sized NO, i.e. N. lui Chou and Ran, 1981 (Chou et al., 1981) and N. carpenteri (Cheng, 1957) indirectly transfer a nuptial gift of a salivary secretion prior to copulation. The short NO of N. lui ends at the base of the PO, and the medium sized NO in N. carpenteri reaches the end of the abdominal segment IV; both stabilize the female wings only partially and apparently play an auxiliary role during copulation. Contrarily, N. longiprocessa (Hua and Chou, 1997), with a long NO capable to cover the entire wing width of the female, exclusively relies on coercive copulation, and does not offer nuptial gifts. Short NOs and POs pinch only the anterior margin of female forewings, and both processes are spatially suited to fit the latter. In Panorpa amurensis MacLachlan, 1872 (MacLachlan, 1872) (Figure 5A,B), the space created by the clasped NO and PO matches the space of the anterior valley fold of the female wing delimited by the convex costal vein (C), the concave subcostal vein (Sc), and the convex radial vein (R₁), clamping together with the PO before or at the latter vein. Our measurements of these distances in this and other panorpid species with various NOs indicate that this seems to be an effective arrangement to grip the anterior wing fold of the female (see Supplementary file 1 for measurements). As the C, Sc, and R₁ run almost parallel over a long stretch, the female forewing may be gripped more anteriorly or posteriorly, usually by its half or basal third, as can be observed in photographs of mating panorpids (Zhong and Hua, 2013a; Zhong et al., 2014; Tong and Hua, 2019).

Figure 5

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A short NO of the type similar to that in Recent panorpids was already present among the Late Jurassic panorpoids, as evidenced by Orthophlebia heidemariae (Willmann and Novokschonov, 1998; Figure 5C, D). This species is currently classified in Orthophlebiidae, a Mesozoic polyphyletic panorpoid assemblage ancestral to Panorpidae (Soszyńska-Maj et al., 2019). In Burmorthophlebia multiprocessa gen. et sp. nov. the NO, although also short, differs from all the hitherto known in Mecoptera, extinct or extant. The NO is distinctly raised over abdomen but poorly pronounced, not reaching any process to form a clamp with. The PO is here replaced by two elevated areas on tergite IV, the postnotal areas 1 and 2 (PA1 and PA2; Figure 5E). The former is mound-like, located after an anterior depression on tergite IV and at a distance from the NO comparable to that of C to R₁. The PA2 is sharply elevated, and measurements show that it contacted the female wing just at or slightly beyond the concave radial sector vein (Rs). Both PAs were involved in adhering the female’s wing as indicated by their location and specialised setation, consisting of short and dense tufts of hairs different from other abdominal bristles. Apparently, only the costal vein and a brief anterior wing membrane portion were covered by the NO in B. multiprocessa gen. et sp. nov., and so the remaining portion of the female wing was to a certain degree immobilised by the setation of both PAs. Among the extant panorpids the hirsute PAs, more flattened than those of Burmorthophlebia gen. et sp. nov. and situated on three consecutive tergites, have been described in Neopanorpa setigera Wang and Hua, 2018 (Wang and Hua, 2018b) and N. luojishana Wang and Hua, 2019 (Wang and Hua, 2019b), and also exist in N. longistipitata Wang and Hua, 2018 (Wang and Hua, 2018b). However, in these species the NO is long and able to keep the entire wing of a female pressed against the male’s abdomen to a lesser or greater extent, so the hirsute PAs are probably only of auxiliary importance (Figure 6A–D). This contrasts with the armature in B. multiprocessa gen. et sp. nov., where it was the interaction between NO-PO setae and the integumentary structures of the female’s wing that kept a larger portion of the latter in place, although the fixing must have been weaker. Recent panorpids with a short NO (and anal horns) adopt a mixed mating strategy (Neff and Svensson, 2013). After a courtship ritual, males present nuptial gifts to the female, but if they are not able to get one ready (including if they have been stolen by other males) or the gift is deemed as unsatisfactory by the female (e.g. small salivary secretions due to poor condition of the male), they resort to attempt coercive copulation (Thornhill and Sauer, 1991). Burmorthophlebia multiprocessa gen. et sp. nov., with its short NO and no other armature to immobilize the female’s wing, probably had a similar approach when mating.

Figure 6

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As introduced above, long NOs differ mechanically from short NOs. They are present in a small group of species classified in the genera Panorpa and Neopanorpa (Hu et al., 2015; Tong and Hua, 2019; Wang and Hua, 2018a). Instead of a pincer-like clamp composed by a PO and NO, the long NO covers the entire female’s wing across its full width and immobilizes it more or less pressed against the abdomen. As the mating couple of N. longistipitata shows (Figure 6A–D), the leading edge of the female wing is also accommodated in a small depression of tergite IV under the base of the NO. As can be deduced from measured distances, of the three hirsute PAs on tergites IV–VI the first one supports the wing fold by the convex R₁; the more distal part of the wing is pressed against the abdomen by the long NO with the help of special muscles (Thornhill, 1990). The tip of the NO is locked between the genital hypovalves. Although this conformation fully encloses the female wing during mating, the resulting grip is likely not particularly tight. In any case, nuptial gifts are not offered prior or during the mating of Neopanorpa species with a long NO, such as N. longiprocessa (Zhong and Hua, 2013a), which would suggest that such process enables highly coercive mating behaviour.

Among extinct mecopterans, an extremely long NO was hitherto only known in Baltipanorpa damzeni from Baltic amber (Krzemiński and Soszyńska-Maj, 2012). In this species, the NO forms together with an unusually elongate PO a clip-like structure able to reach beyond the full width of the female’s wing during mating (Figure 6E). Both NO and PO possess short tooth-like bristles, which would have increased their holding power. The PO is bent anteriorly, partially securing the clip in this way. Baltipanorpa oppressiva sp. nov has a similarly characteristic NO-PO clip bearing inner tooth-like bristles, but its NO is terminally forked in two prongs bent posteriorly at an almost right angle, which embrace the tip of PO (Figures 1B, E, 6F). The interaction of these structures in the new species results in a full clasp-like mechanism. The female wing enclosed in this remarkable clamp was likely fully immobilised, a withdrawal attempt probably causing risk of tearing the wing membrane. A similar function, although less effective than that in B. oppressiva sp. nov, must have been accomplished in B. damzeni by a few pairs of elongate bristles on the posterior side of the NO (Figure 6E). Furthermore, the NO-PO clamp in Baltipanorpa is raised at angle of about 50° from the abdomen, and so the captured female’s forewing was also kept raised and separated from the male’s abdomen. This position is deemed as genuine (not preservational) as it is now known from two different species (Figure 6E and F) and is unique to this genus. In extant panorpids with very long NOs (Neopanorpa), this process is placed at a much lower angle and keeps the female wing rather flat against the abdomen, having no other counterpart such as the long PO in the two fossil Baltipanorpa species discussed herein (Figure 6A and D).

Conclusions

The morphology of the male’s abdomen in extant scorpionflies represents a good approximation to the mating strategy used toward a female. Thus, the abdominal structures of fossil specimens have the ability to convey information on the likely reproductive strategies of extinct scorpionflies and, to some extent, on their evolution. Based on the disparate male abdominal shapes of the three fossil taxa described herein, these almost certainly had different mating strategies along a nuptial gifting-coercive gradient as observed among their extant panorpid relatives. Firstly, the Cretaceous species Cantabra soplao gen. et sp. nov., currently the only known member of its lineage, bears a small PO, with the absence of other posterior abdominal processes. This suggests that its mating behaviour was devoid of coercion and that nuptial gifting must have played an important role in these scorpionflies. Extant panorpid species which NO-PO system is very small or even absent and lack posterior abdominal processes (‘horns’) either rely on nuptial edible gifts or on mouth-to-mouth feeding during copulation adopting an O-shaped position instead of the typical V-position (Figure 6A; Zhong et al., 2015). It is plausible that the mating in C. soplao gen. et sp. nov. was similar. Regardless, the great degree of elongation of the abdomen in this species was probably involved in sexual selection dynamics.

Secondly, the Cretaceous Burmorthophlebia multiprocessa gen. et sp. nov., the only orthophlebiid specimen described from amber to date, likely exhibited a mixed mating strategy involving both nuptial gifting and coercive behaviour, as observed in some extant Panorpidae species. This inference is based on a weak notal-postnotal clamping system and two sets of paired processes on the abdomen, which likely ensured female restraint and probably extended copulation beyond any possible nuptial gift stages. Noteworthy, abdominal processes are exceptionally abundant in this species, on the dorsal and ventral sides. Such armature is unknown among extant Panorpidae, and can be compared only with that of Nothiotrauma reedi of the relict family Eomeropidae.

Lastly, in the Eocene Baltipanorpa oppressiva sp. nov. the NO-PO wing clamp, which resembled a sealing clip, was able to tightly grasp the full wing width of the female. Unlike its previously known congeneric relative B. damzeni, this structure bore a sophisticated terminal clasp. Together with a highly recurved abdominal horn in this new species, these traits represent the most extreme set of female clamping devices known among scorpionflies, both extinct or extant. Consequently, a fully coercive mating strategy was likely dominant, even exclusive, in this species. During copulation, the female wings (at least the forewings) were likely kept in the notal-postnotal clamp well raised over the male abdomen, this way being separated from the male’s anal horn, which is positioned more proximally in this species (i.e. on tergite V vs. in tergite VI in other panorpids). This inclined position of the female’s wing(s) enabled the entrapment of her abdomen by the male’s anal horn during mating. In evolutionary terms, the fitness costs of such organs and the resulting coercive mating behaviour might have been too high for Baltipanorpa. The very long notal and postnotal processes (Figure 6E and F) raised over the male’s body might have hindered flight ability; moreover, because of their size (and perhaps their permanently raised position) these processes were likely prone to injury and thus they might entail a risk for their bearers. These factors might have contributed to the extinction of Baltipanorpa and its singular morphology, so that the evolution of this extreme clamping device resulted in a blind alley. Moreover, the biology of the female could have also contributed to this extinction if the mating became too oppressive or dangerous, related to the risk of tearing the wing membrane with the teeth and hard bristles of the male’s notal organ.

The remarkable diversity of abdominal configurations shown herein suggests that panorpoid mecopterans had accordingly disparate mating strategies and behaviours in the past. Data based on fossil and extant species suggest that the coercive mating behaviour in this group is secondary (derived) from the habit of nuptial gifting. According to the current fossil record, long notal organs did not evolve prior to the Eocene (Figure 7; Supplementary file 2). Data from extant mecopterans point at the Choristidae as the sister group of Panorpoidea (Willmann, 1989; Wang and Hua, 2021), which lack notal organs or other abdominal modifications (including elongate distal segments) and adopt an O-shaped mating position while the male transfers mouth-to mouth salivary secretion to the female (Zhong et al., 2015). Hence, it is assumed that the latter condition is likely plesiotypic for Panorpoidea.

Figure 7

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Our data provide first steps toward gaining a deep-time perspective able to inform discussions on the evolution of mating-related sexual conflict in Mecoptera and in insects more generally. Extant fauna show that extreme female holding mechanisms such as those from Baltipanorpa did not survive until today among scorpionflies (Wang, 2020). In fact, mating strategies involving gift provisioning appear to have been evolutionarily favored among mecopterans contrary to forced mating, and likely more prone to promote the species' genetic diversity. In order to start formulating hypotheses about when mating strategies –including those fully coercive– and associated anatomical structures of scorpionflies appeared throughout evolutionary history, it will be necessary to keep discovering fossil males with good abdominal preservation. The latter will in turn necessitate from improving the knowledge on the currently contentious phylogenetic relationships among mecopteran groups.

All data needed to evaluate the conclusions in the paper are present in the paper and/or the supplementary materials. Additional information related to this paper may be requested from the authors. Investigated fossils are available in public institutions: at the Institutional Collection from the El Soplao Cave (Government of Cantabria), Celis, Cantabria, N Spain and at the collection from the Museum of the Institute of Systematics and Evolution of Animals (ISEA), Polish Academy of Sciences (PAS), Kraków, Poland.

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Author details

1. Agnieszka Soszyńska-Maj

   Faculty of Biology and Environmental Protection, Department of Invertebrate Zoology and Hydrobiology, University of Łódź, Łódź, Poland

   Contribution

   Conceptualization, Investigation, Methodology, Resources, Software, Validation, Visualization, Writing – original draft, Writing – review and editing

   For correspondence

   agnieszka.soszynska@biol.uni.lodz.pl

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-2661-6685

2. Ewa Krzemińska

   Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland

   Contribution

   Conceptualization, Validation, Visualization, Writing – original draft, Writing – review and editing

   For correspondence

   ekrzeminska9@gmail.com

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-3431-9963

3. Ricardo Pérez-de la Fuente

   Oxford University Museum of Natural History, Oxford, United Kingdom

   Contribution

   Conceptualization, Investigation, Resources, Validation, Visualization, Writing – original draft, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-2830-2639

4. Ji-Shen Wang

   College of Agriculture and Biological Sciences, Dali University, Dali, China

   Contribution

   Investigation, Methodology, Resources, Validation, Visualization

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-0188-0228

5. Krzysztof Szpila

   Department of Ecology and Biogeography, Faculty of Biological and Veterinary Sciences, Nicolaus Copernicus University, Toruń, Poland

   Contribution

   Conceptualization, Validation, Writing – original draft, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-3039-3146

6. Kornelia Skibińska

   Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland

   Contribution

   Investigation, Software, Validation, Visualization

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-5971-9373

7. Katarzyna Kopeć

   Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland

   Contribution

   Data curation, Investigation, Project administration, Resources, Validation

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-6449-3412

8. Wiesław Krzemiński

   Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland

   Contribution

   Conceptualization, Funding acquisition, Investigation, Resources, Supervision, Validation, Writing – original draft, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-5685-891X

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